JOURNAL OF FOREST SCIENCE, 49, 2003 (5): 208–219
Composition of psocid taxocenoses (Insecta: Psocoptera) in dependence on the level of naturalness of forest ecosystems in the Žďárské vrchy hills P. MÜCKSTEIN1, O. HOLUŠA2 Masaryk University, Faculty of Science, Brno, Czech Republic Mendel University of Agriculture and Forestry, Faculty of Forestry and Wood Technology, Brno, Czech Republic 1 2
ABSTRACT: In 1999–2000 in the Protected Landscape Area (PLA) Žďárské vrchy hills the occurrence of psocids (Psocoptera) was studied in different types of biotopes: natural forest ecosystems (stands of Fagus sylvatica with individual admixture of Abies alba, Acer pseudoplatanus, Picea abies), changed forest ecosystems (monoculture of Picea abies), young plantations in forest stands, disperse forest vegetation (solitary trees), and also non-forest ecosystems – agrocenoses, meadows and grazing lands. A total of 10,560 adults in 20 species were found. Three groups of biotopes with specific psocid taxocenosis were found out by computed cluster analysis – l. natural forest stands with dominance of Fagus sylvatica, 2. disperse tree vegetation and solitary trees in cultural landscape, and 3. forest stands remote to nature (monoculture of Picea abies). Occurrence of psocids was observed from the beginning of May to mid- November. Maximum of abundance was found in September. Keywords: Psocoptera; types of forest ecosystems; Protected Landscape Area Žďárské vrchy; Žďárský biogeographical region; Czech Republic
In general, psocids are a rarely studied insect order. Thank to their size, quiet coloration and relatively difficult way of collection and preparation they are on the margin of entomologists’ interest. In the Czech Republic the psocids were studied only in some areas [mostly in the mountains of Moravia and Silesia – Hrubý Jeseník Mts., Králický Sněžník Mts. (OBR 1949), and in the Moravskoslezské Beskydy Mts. (OBR 1952, 1965)]. Only occasional collections have been reported from other areas. Sporadic faunistic data from the area of the Žďárské vrchy hills were presented by OBR (1948, 1958). No complex psocopterological research in this area has been carried out. At present, only faunistic data are mostly known from this country; HOLUŠA (2001) studied an ecological problem of the composition of psocid taxocenoses in dependence on vegetation zones in the area of the Moravskoslezské Beskydy Mts. The aim of this paper is to evaluate the occurrence of psocids in various types of biotopes in the Žďárské vrchy hills. We attempted to find out the psocid taxocenosis composition in dependence on the level of biotope naturalness or to find out the influence of the growth period of forest
ecosystems on the composition of psocid taxocenoses. In this paper results of two-year faunistic-ecological research on the Psocoptera order that was carried out in selected localities of the PLA Žďárské vrchy are presented. METHODS Research was carried out in 1999–2000. In the studied area permanent sites were marked out in various types of biotopes so that as high as possible variety of ecological conditions would be registered. The material was collected at permanent sites (Table 1) during the vegetation period (from the beginning of May to mid-November) in 5 day intervals. The material was obtained by sweeping using a sweeping net of 50cm frame diameter. Branches of trees and bushes were beat with the same sweeping net at a distance of about 1 m from the branch end and up to about 2.5 m of height. These methods were also complemented by individual collecting of adults. When sweeping and beating, 30 sweepings or beatings were done in particular localities. Caught psoc-
This paper was worked out in the framework of Research Project CEZ J 8/98:434100005 of Faculty of Forestry and Wood Technology of Mendel University of Agriculture and Forestry Brno.
208
J. FOR. SCI., 49, 2003 (5): 208–219
Altitude (m)
Altitudinal vegetation zone
Exposure
Type of biotope
Level of naturalness
Description of tree species composition (%)
Žákova hora
6361
802
5.
JV
1
2
fs 80, pa 10, ap 5–8, aa 2–5
Ranský Babylon
6360
670
5.
SV–V
1
2
fs 60, pa 40
6362 6362
828 823
5. 5.
JV S–0
2 2
4 4
pa 100 pa 100
6362
650
5.
SV
2
4
pa 100
6361 6261
703 654
5. 5.
Z–JZ SZ
2 2
4 4
Local name of locality
Code of grid mapping
ZKH
Cadastral territory
Abbreviation of locality
Table 1. Description of places of psocid collection
DSK KJV
Cikháj Staré Ransko Křižánky Fryšava
KR1
Křižánky
CKJ HL1
Cikháj Hlinsko
Devět skal Křivý javor Křižánecké polesí U Snítku Pláňavy
HER
Herálec
Na Čermačkách
6361
590
5.
J–JV
3
5
KR2
České Křižánky
6362
708
5.
Z
3
5
Ranské polesí
6360
620
5.
S
3
5
pa 80, ps 10, fs 10
MIL
Křižánky Staré Ransko Milovy
pa 100 pa 100 pa 30, bv30, ap 20, fs 10, ca 10 pa 90, fs 10
Milovské polesí
6362
721
5.
SV
3
5
HL2
Hlinsko
Čertovina
6261
673
5.
J–JZ
4
5
JIM
Jimramov
Holý vrch
6363
583
5.
V
4
5
pa 70, ld 30 tc 40, pa 30, ah 10, bv 10, ca 5, qr 5, ms+ qr 10, pa 30, ps 10, ld 10, tc 25, fe 10, ca 5
6462
628
5.
SV–V
4
5
pa 50, sa 20, tc 20, fs 10
6361
652
5.
V
4
5
pa 60, ap 20, fs 10, ps+
6261
585
5.
S–0
4
5
6362 6363 6361
660–690 500–550 690–700
5. 5. 5.
SV–V Z–J JZ
5 5 5
5 5 5
ps 20, pa 30, bv 20, psp 10, tc 10, ms+ – – –
6361
570
5.
Z
6
6
–
6261
600
5.
V
6
6
–
RBB
STR
NMM STV KAM KDL JIL CKA ZSA HLA
Nové Město Harusův kopec na Moravě w-shore of Strž Světnov dam Vojtěchův Kameničky kopec Kadov – Jimramov – Cikháj municipality Žďár nad Tálský mlýn Sázavou Hlinsko Blatno
ids were sucked with an exhauster and stored in a small test tube with 70% of alcohol. The material was collected and determined by P. MÜCKSTEIN. Evidence material is deposited in 70% of alcohol in the collection of the first author of this paper. Papers of GÜNTHER (1974) and LIENHARD (1998) were used for determination; nomenclature, zoogeographical distribution and ecological demands are according to LIENHARD (1998). When the conditions of biogeocenoses are described, specific names of forest growth phases are introduced – clearcut area, young plantation, pole timber stand, polestage, large-diameter stand (according to KOLEKTIV 1994, 1995). Classification of the level of naturalness was done according to ELLENBERG (1973, 1978). Classification into units of forest typology (PLÍVA 1971, 1991) was carried out on the basis of our own investigations. Values of dominance, species diversity and equitability were computed from obtained psocid numbers. ShannonWiener and Simpson’s indices were used for the calculation J. FOR. SCI., 49, 2003 (5): 208–219
of species diversity at particular localities. Faunistic similarity was calculated according to Jaccard’s and Sörensen’s similarity indices. For a comparison of the localities Renkonen’s similarity index was also used that originates from the locality comparison on the basis of the recorded species dominance. Cluster analysis was computed by help of the programme Statistics for Windows 5.5. For calculation the Euclidean distances and methods Complete Linkage, Unweighted Pair Group and Average and Ward’s method were used. Collection of the material in the PLA Žďárské vrchy was permitted by the exception 14697/98-OOP/5073/98-V408 of the Ministry of Environment. In Table 1 these abbreviations of tree species names are used: aa – Abies alba, ah – Aesculus hippocastanum, ap – Acer pseudoplatanus, bv – Betula verrucosa, ca – Corylus avellana, fe – Fraxinus excelsior, fs – Fagus sylvatica, ld – Larix decidua, ms – Malus sylvestris, pa – Picea abies, ps – Pinus sylvestris, psp – Prunus spinosa, qr – Quercus robur, sa – Sorbus aucuparia, tc – Tilia cordata. 209
Study area All localities where research on psocid taxocenoses was carried out are situated in the PLA Žďárské vrchy (Fig. 5) that lies in the north-eastern part of the Českomoravská vrchovina Highlands. In accordance with biogeographical classification the whole studied area belongs to the Žďárský biogeographical region (1.65) (cf. CULEK 1996). The bioregion has a character of flat or broken highland with quite narrow peak parts, valleys are deep and broad. The whole area lies in the zone “oreophyticum”. Vegetation has a submontane or montane character. The 5th firbeech (Abieto-Fagetum) vegetation zone predominates, the 4th beech (Fagetum) vegetation zone is represented partly, azonal communities of natural pinewoods occur sporadically on Histosols (according to the systematics of PLÍVA 1971, 1991). In the biogeographical region submontane forest fauna of the Hercynian origin predominates; it is relatively best preserved in rests of original beech growths. At present peat-bog fauna, which is fairly marked, is recedent because of drainage. Climatically the area belongs to the cold region CH7 (QUITT 1975) with the following climatic characteristics: number of days above 10ºC: 120–140; mean temperature in January: –3 to –4ºC; mean temperature in July 15–16ºC; mean temperature in April: 4–6ºC; mean temperature in October: 6–7ºC; precipitation amount during the vegetation period: 500–600 mm; precipitation amount in winter: 350–400 mm. Only marginal parts of the area lie in the contact with moderately warm region MT 3. Description of localities Types of ecosystems 1. Natural forest stands and near-natural stands (level of naturalness 2 according to ELLENBERG 1973, 1978) – forest stands with well-preserved tree composition that is constituted by dominant European beech (Fagus sylvatica), with admixture of Norway spruce (Picea abies) while silver fir (Abies alba) and sycamore maple (Acer
pseudoplatanus) occur sporadically. Stands have a vertical canopy at some places. The species Asperula odorata, Brachypodium sylvaticum, Calamagrostis arundinacea, Majathemum bifolium, Senecio ovatus dominate in the undergrowth; at present only remnants of this type of biotope are preserved at higher locations of the Žďárské vrchy. Classification into units of forest type system: 5S (Abieto-Fagetum mesotrophicum) and 5B (Abieto-Fagetum eutrophicum). 2. Forest stands remote to nature (level of naturalness 4) – forest stands (monocultures) of Picea excelsa, only individually the admixture of Fagus sylvatica, evenaged stands with horizontal canopy. Undergrowth is poor and consists of several species only, in younger stands it is missing: Oxalis acetosela, Majanthemum bifolium, Deschampsia flexuosa, Luzula nemorosa, Calamagrostis villosa. Classification into units of forest type system: 5S (Abieto-Fagetum mesotrophicum), 5H (Abieto-Fagetum illimerosum trophicum), partly 5O ((Fageto)-Abietum variohumidum trophicum). 3. Young plantations with clearcut vegetation in forest stands (level of naturalness 5) – clearcut areas of Picea abies, with admixture of Larix decidua and Pinus sylvestris. Dominant species of vegetation under trees – Rubus idaeus, Chamaenerion angustifolium, Calamagrostis epigeois. Adjacent stands are constituted by pole timber stages or large-diameter stands of Picea abies. 4. Disperse forest vegetation and solitary trees (level of naturalness 5) – solitary old trees (Quercus robur, Tilia cordata, Acer platanoides, Acer pseudoplatanus), small groups of pole timber stage and pole-stage stands of Betula verrucosa, Sorbus aucuparia, Cerasus avium, Corylus avellana, Prunus spinosa, individually the admixture of Fagus sylvatica. 5. Meadows (level of naturalness 5) – intensively cultivated and fertilised grasslands. Vegetation cover is constituted by the species: Festuca rubra, Trifolium repens, Cynosorus cristatus, Artemisia vulgaris, at places the growth of Nardus stricta. 6. Agrocenoses (level of naturalness 6) – arable land with intensive production of agricultural crops. Collection was
Number of adults 1,800 1,600 1,400 1,200
1999
1,000
2000
800 600 400 200 0 V.
210
VI.
VII.
VIII.
IX.
X.
XI.
Month
Fig. 1. Seasonal dynamics of psocids in the Protected Landscape Area Žďárské vrchy in 1999 and 2000 (number of adults)
J. FOR. SCI., 49, 2003 (5): 208–219
carried out in one-year monocultures of oat (Avena sativa) or rye (Secale cereale). RESULTS AND DISCUSSION When investigating selected localities in the area of the PLA Žďárské vrchy during the vegetation periods in 1999 and 2000, a total of 10,560 specimens of free living psocid imagoes in 20 species were obtained that belong to 6 families of the suborder Psocomorpha. Out of the total species number of the suborder Psocomorpha living in the Czech Republic it is about 37%. It was possible to plot a graph of psocid seasonal dynamics in 1999 and 2000 on the basis of specimen collections at regular time intervals (about 5 days) (Fig. 1). The first imagoes were recorded already in the first decade of May (Loensia fasciata), the last imagoes were caught in mid-November (Caecilius burmeisteri). Maximum abundance in both years was recorded in September. The localities where psocids were collected were selected in such a way that their ecological conditions would be different if possible. Out of the original 21 selected localities (Fig. 5, Table 1) the psocids were not recorded at all or they were recorded at an amount of only several specimens in 5 localities CKA, ZSA, HLA, KDL and JIL. Although there exist literary data on psocids inhabiting also the above-mentioned biotopes (OBR 1948, 1951a,b, 1958, 1959), results from these localities were more or less negative. Only in the agrocenosis in CKA locality 4 imagoes of Caecilius burmeisteri and 2 imagoes of Caecilius flavidus in KDA locality were found. But these species probably originated from a nearby forest stand of Picea abies, from where they could be blown away. During the experimental psocid collection into a light collector in the village of Herálec (faunistic square code 6361, garden, 653 m above sea level, July 1999) a mass arrival of the species Metylophorus nebulosus (364 imagoes) was recorded.
It results from the recorded psocid species spectrum in the area of the Žďárské vrchy that the given area is inhabited by the most common and resistant psocid species with greater occurrence range in the vegetation zones. From the total number the species Philotarsus picicornis, Caecilius burmeisteri, Caecilius flavidus and Loensia fasciata were eudominant. The species composition is probably influenced by a relatively high altitude of the localities, rather cold climate and relatively poor and monotonous composition of vegetation (dominance of Picea abies monocultures). The species typical of higher altitudes (Metylophorus nebulosus, Trichadenotecnum majus, Amphigerontia bifasciata), and also species preferring lower altitudes (Graphopsocus cruciatus, Caecilius flavidus) correspond with the character of the 5th (fir-beech) vegetation zone (Abieto-Fagetum). Cluster analysis In cluster analysis dendrograms were computed by help of three methods. Complete Linkage (Fig. 2) and Unweighted Pair-Group Average methods (Fig. 3) provided approximately identical results. These methods divided the localities into two main groups, and either of them is divided into two subgroups. In the first main group the a-subgroup is represented by two localities – RBB and ZKH, i.e. localities of natural character. Within the first main group the b-subgroup comprised localities KAM, JIM, HL2, NMM and STV is associated with the a-subgroup. These localities are represented by the biotopes of solitary old trees (Quercus robur, Tilia cordata, Acer pseudoplatanus), at some places surrounded by small groups of Betula verrucosa, Sorbus aucuparia, Cerasus avium and Corylus avellana (character of young growth stand or small pole stage). The second main group is represented by the a-subgroup with localities DSK, STR, HER, MIL, KR2, KR1, CKJ, HL1, remote localities to natural stands, with Picea abies dominance. The b-sub-
TreeDiagram Diagram 16 Variables Tree for 16for Variables Complete Linkage Complete Linkage
Euclidean distances Euclidean Distance ZKH RBB KAM JIM HL2 NMM STV DSK STR HER MIL KR2 KR1 CKJ HL1 KJV
00
50 50
100 100
150 150
Linkage Distance Linkage Distance
J. FOR. SCI., 49, 2003 (5): 208–219
200 200
250 250
Fig. 2. Dendrogram of mutual similarity of localities according to Renkonen’s index (used cluster methods Complete Linkage and Euclidean Distances)
211
Tree Diagram for 16 Variables Tree DiagramPair-Group for 16 Variables Unweighted Average Unweighted pair-group average Euclidean Distance
Fig. 3. Dendrogram of mutual similarity of localities according to Renkonen’s index (used cluster methods Unweighted Pair-Group Average and Euclidean Distances)
Euclidean distances
ZKH RBB KAM JIM HL2 NMM STV DSK STR HER MIL KR2 KR1 CKJ HL1 KJV
0
50 50
100 100
150 150
200 200
Linkage Distance
Linkage Distance
Tree Diagram for 16 Variables Tree Diagram for 16 Variables Ward’smethod method Ward`s Euclidean Distance Euclidean distances ZKH RBB KAM JIM HL2 STV NMM DSK STR HER MIL KR2 KJV KR1 CKJ HL1
00
200 200
400 400
600 600
Linkage Distance
Linkage Distance
group with the only one locality KJV is associated with previous subgroup. A dendrogram computed by Ward’s method (Fig. 4) was different: the second main group was divided into the a-subgroup with localities DSK, STR, HER, MIL, KR2 (localities with young forest plantation of Picea abies with clearcut vegetation in complexes of large-diameter stands and pole-stage stands of Picea abies; Larix decidua (Mill.) was an admixed species at some places in the young plantation only) and the b-subgroup with localities KJV KR1, CKJ, HL1, localities of the biotope of Picea abies monoculture. Division of the first main group and its subgroups was the same as it was in the case of previous methods. It is possible to see from the dendrograms (Figs. 2–4) that the 16 studied localities were divided approximately into three different groups – 1. psocid taxocenoses living in forest stands with dominance of Fagus sylvatica with naturalness level 2; 2. taxocenoses inhabiting dispersed and solitary vegetation in open cultural landscape; 3. psocid taxocenoses in monocultures of Picea abies. However, the 212
800 800
1,000 1000
Fig. 4. Dendrogram of mutual similarity of localities according to Renkonen’s index (used cluster method Ward’s method and Euclidean Distances)
types of biotopes 1 and 2 are closer to each other that the biotopes with dominance of deciduous trees contrary to the type of biotope 3, which is coniferous communities. 1. Forest stands with dominance of Fagus sylvatica: psocid taxocenoses in localities ZKH and RBB are characterised by the presence of folicolous and corticolous species. In both localities the eudominant species are Caecilius flavidus, Philotarsus picicornis, Loensia and Stenopsocus lachlani. In RBB locality the species Caecilius burmeisteri was furthermore recorded as eudominant, but it was not recorded in locality ZKH. Elipsocus pumilis was a dominant species for both localities and Metylophorus nebulosus was a recedent species. The finding of the psocid Psococerastis gibbosa in RBB locality is surprising. This non-abundant species is typical of deciduous stands in open landscape. On the whole, 10 species of psocids were found in this type of biotope (ZKH – 7 species, RBB – 9 species) (Table 2). 2. Dispersed vegetation and solitary trees in cultural landscape: psocid taxocenoses were represented by the
J. FOR. SCI., 49, 2003 (5): 208–219
J. FOR. SCI., 49, 2003 (5): 208–219
213
10.9
10.0
3.6
8.4
28.3
CKJ
8.3
0.6
4.9
4.4
15.9
HER
11.7
12.7
0.8
Loensia fasciata
Loensia variegata
7
23.15
2.31
0.82
0.76
Total number of species
Abundance
Shannon-Wiener index of diversity
Equitability
Simpson index of diversity
0.80
0.83
2.62
29.35
0.88
0.94
3.25
21.6
11
432
9
11.6
463
Total number of specimens
4.2
Trichadenotecnum majus 587
0.2
Psococerastis gibbosa
2.5
0.74
0.75
2.39
24.7
9
495
5.2
20.4
0.81
0.86
2.71
45.5
9
911
3.7
6.7
0.84
0.85
3.06
27.65
12
553
4.7
1.1
6.5
0.86
0.92
2.13
40.51
10
811
7.2
10.9
0.88
0.91
3.27
32.12
12
642
9.9
3.6
4.9
6.7
0.88
0.92
3.31
31.35
12
627
10.5
2.9
4.5
10.0
0.87
0.93
3.22
42.55
11
811
10.2
11.6
12.0
7.0
0.89
0.87
3.48
76.55
16
1,531
1.7
7.2
6.8
4.9
12.1
2.8
7.9 10.3
Metylophorus nebulosus
1.6
2.3
16.5
5.9
6.6
3.8
8.8
17.4
9.1
0.6
7.2
16.9
5.4
2.5
1.1
8.5
6.5
19.9
STR
3.5
16.0
8.0
1.9
4.3
4.9
13.5
17.8
KR2
Amphigerontia bifasciata
18.9
11.1
4.5
4.5
0.5
4.5
14.0
18.2
MIL
Mesopsocus unipunctatus
31.3
17.3
3.6
0.6
6.1
9.8
10.4
7.8
23.2
HL1
0.5
42.8
12.5
6.2
1.8
20.0
KR1
Mesopsocus laticeps
16.7
25.7
Philotarsus picicornis
29.6
6.9
Philotarsus parviceps
2.0
6.6
2.2
1.4
1.6
2.6
11.3
KJV
11.4
Reuterella helvimacula
9.1
7.2
2.5
17.4
DSK
Elipsocus pumilis
11.2
28.2
11.5
RBB
0.7
12.1
32.9
ZKH
Locality
Cuneopalpus cyanops
Stenopsocus lachlani
Graphopsocus cruciatus
Enderleinella obsoleta
Caecilius piceus
Caecilius flavidus
Caecilius despaxi
Caecilius burmeisteri
Species
Table 2. Dominance of psocids and structural features of psocid taxocenoses at the places of collection
0.85
0.86
2.98
33.91
11
678
5.5
0.6
22.3
0.3
1.6
13.6
9.7
7.2
9.6
20.4
9.3
KAM
0.84
0.83
2.88
41.25
11
825
8.4
0.5
22.0
0.6
0.7
13.2
11.0
6.3
11.5
23.5
2.3
JIM
0.84
0.92
2.76
21.31
8
426
16.4
16.9
0.5
13.1
11.5
6.1
22.1
13.4
STV
0.76
0.87
2.24
13.81
6
276
1.2
20.3
15.2
9.8
36.5
17.0
NMM
0.78
0.81
2.67
24.61
10
492
8.7
0.2
23.3
0.2
1.0
18.2
7.9
7.5
12.1
26.6
HL2
44.4
45.4
40.0
42.1
62.5 77.8 95.2
70.5 70.5
85.7 38.4
27.2 35.3
38.1 45.4
33.3 33.3
36.3 20.0
12.5
47.6
31.5
47.1
13.1 43.4 HL2
NMM
61.5
53.3
26.6
100
45.4 100 66.6
63.6 55.6 100 33.3
50.0
50.0
52.6
50.0
84.2
84.2 60.0 58.8 63.1 STV
53.3
70.5
47.1
90.9
90.9
54.5 72.7 100
51.8
51.8 45.5
45.5 43.4
43.4 43.7
43.7 28.5
28.5
54.5 72.7 100
52.1 50.0
100
100 52.1 50.0
54.5
40.0
54.5
60.0 52.6 JIM
KAM
52.6
60.0
40.0
31.3
23.5 35.7 29.4
33.3 81.4 85.7
86.9 86.9
85.7 76.9
76.1
22.2 35.0 100 85.7 72.0
21.4
28.6
29.4 68.8 100 86.9 80.0
81.4
64.0
90.9
56.0 43.4 HER
STR
55.5
60.0
70.0
22.2
22.2 33.3
33.3 27.8
27.8 27.8 75.0
27.8 91.6 81.8
81.8
20.0 76.9 100 91.6 76.2
20.0
75.0
76.9 84.6 100 83.3 76.2
86.9
66.6
86.9
57.1 42.1 KR2
MIL
42.1
47.6
76.1
61.5
62.5
16.7
16.7
20.0
6.7
11.1 69.2 69.2 100 72.7 63.1 66.6 HL1
35.2
52.6
84.2
29.4
26.7 41.7
42.9 35.3
33.3 33.3
35.3 75.0
56.3 66.7
76.9 84.6
61.5 61.5
28.6 57.1 100
25.0
71.4
46.1 75.0 100
85.7 95.6
57.1
90.0
57.1 52.6 CKJ
KR1
33.3
44.4
55.5
31.3
18.8 30.8
46.2 37.5
25.0 25.0
37.5 68.8
47.1 53.8
83.3 76.9
50.0 61.5
76.9 50.0
15.4 40.0 38.4
30.8
72.7
91.6 81.8
100 55.5
42.8 100
60.0
50.0
33.3
44.4 DSK
KJV
35.7
41.6 36.3
54.5 42.8
38.4 38.4
42.8 38.8
27.7 28.5
42.8 40.0
26.6 26.6
31.2 35.7 33.3 100
21.4
38.4
28.6 60.0
75.0
100 ZKH
RBB
33.3
36.3 40.0 28.5
44.4 26.6 23.1
HL2 HL1
MIL
KR2
STR
HER
KAM
JIM
STV
NMM CKJ KR1 KJV DSK RBB ZKH
Table 3. Values of Jaccard’s and Sörensen’s indices of similarity for particular localities
214
species that are typical of sites with lower altitude. Eudominant species in these biotopes are: Caecilius flavidus, Loensia fasciata a Mesopsocus unipunctatus. The species Graphopsocus cruciatus, Trichadenotecnum majus, Mesopsocus laticeps and Elipsocus pumilis are dominant. In total 9 species of psocids were recorded in this type of biotope (Table 2). 3. Monocultures of Picea abies: these psocid taxocenoses are characterised by high abundance. Eudominant species are: Philotarsus picicornis, Caecilius burmeisteri and Metylophorus nebulosus. In total 16 species of psocids were recorded in this type of biotope. Psocid taxocenoses in particular localities are more or less similar to each other, only HER locality, where 16 species were found, differs markedly. A relatively high number of species found in this locality can be explained by the south-eastern exposition of this site, also protected from wind. For this reason the microclimate of this locality is different from that of other localities. The finding of Cuneopalpus cyanops, which was recorded only in this locality, is noticeable. Cluster analysis pointed out that psocid taxocenoses inhabiting biotopes of forest clearcut areas do not differ very much from those living in pole-stage stands and large-diameter stands of Picea abies. The species composition of psocids on newly reforested clearcuts (young plantations of Picea abies or Pinus sylvestris) do not anywise differ from large-diameter stands of Picea abies monocultures. Characteristic species of this biotope are: Amphigerontia bifasciata, Philotarsus picicornis, Philotarsus parviceps, Caecilius burmeisteri, C. despaxi, C. piceus, Metylophorus nebulosus, Trichadenotecnum majus, Loensia fasciata, Elipsocus pumilis, Stenopsocus lachlani, Mesopsocus unipunctatus and Reuterella helvimacula. In total 16 species of psocids were recorded in this type of biotope (Table 2). Diversity The values of Shannon-Wiener index ranged from 2.13 to 3.48. Equitability values were in the range from 0.75 to 0.94. In the localities with naturalness level 2 the values of this index were 2.31 to 2.62 (equitability 0.82–0.83), in the localities with naturalness 4 level these values were 2.13–3.25 (0.75–0.94) and in the localities with naturalness level 5 the values were in the range of 2.24–3.48 (0.81–0.93) (Table 2). List of species Caeciliusidae Caecilius burmeisteri Brauer, 1876 Folicolous species living on deciduous and coniferous trees. In the PLA Žďárské vrchy it
J. FOR. SCI., 49, 2003 (5): 208–219
NMM
HL2
0.68
100
100
STV 0.66 0.70 100
0.93
0.87 0.71
JIM 0.69 100
KAM
0.66
0.72 0.90 100
0.24
0.23 0.30
0.40 0.42
0.48 0.25
MIL
HER 0.34 100
100
0.19
0.21 0.29
0.27 0.38
0.70 0.23
0.21 0.26
0.28 0.73
STR
0.25
0.30 0.71
0.75 100
0.08 0.17 0.32 0.10 0.16 0.63 0.76
KR2 0.74
0.75 0.96 100
HL1 0.69 0.68 100
0.20
0.16 0.30
0.32 0.39
0.34 0.17
0.22 0.30
0.26 0.72
0.68 0.62
0.69 0.69
0.61 0.62
CKJ 0.70 100
KR1
0.66
0.56 0.72 100
0.24
0.09 0.14
0.34 0.51
0.32 0.11
0.25 0.31
0.19 0.50
0.78 0.84
0.55 0.48
0.75 0.75
0.47 0.57
0.68 0.73
KJV 0.66 100
0.46
0.45 0.58
0.56 0.44
0.54 0.45
0.44 0.42
0.50 0.56
0.47 0.33
0.44 0.41
0.27 0.28
0.39 0.37
0.25 0.32
0.43 0.46
0.39 0.37
DSK
0.54
0.67 0.54 100
100
RBB 0.42
0.86 100
DSK
KJV
KR1
CKJ
HL1
MIL
KR2
STR
HER
KAM
JIM
STV
NMM
HL2
ZKH
0.44
0.33
RBB ZKH
Table 4. Values of Rekonen’s index of similarity
J. FOR. SCI., 49, 2003 (5): 208–219
was recorded in 14 localities. Eudominant species within the whole studied area. C. burmeisteri has two generations per year in Central Europe, it overwinters in the egg stage. Distribution: Holarctic. Caecilius despaxi Badonnel, 1936 Folicolous species occurring predominantly on coniferous trees. In the PLA Žďárské vrchy all specimens were caught exclusively on spruce (Picea excelsa). Subdominant species within the whole studied area. Distribution: Palearctic. Caecilius flavidus (Stephens, 1836) Folicolous species living more often on deciduous trees. Only parthenogenetic females occur in Central Europe. Males, which are very rare, were found only in Switzerland (LIENHARD 1977). It has as many as three generations per year. Eudominant species within the whole studied area. Distribution: Holarctic. Caecilius piceus Kolbe, 1882 Folicolous species, most frequently on coniferous trees (often together with C. burmeisteri). In Central Europe it has two generations per year. Subdominant species within the whole studied area. Distribution: western Palearctic. Enderleinella obsoleta (Stephens,1836) Folicolous species often occurring on spruce together with C. burmeisteri, but always in smaller populations. It has two generations yearly in Central Europe. Subrecedent within the whole studied area. Distribution: western Palearctic. Stenopsocidae Graphopsocus cruciatus (Linnaeus, 1768) Folicolous species living on deciduous trees. In the PLA Žďárské vrchy found only on solitary deciduous trees (Quercus robur, Malus sylvestris). Subdominant species within the whole studied area. Distribution: Palearctic (LIENHARD 1998), introduced into North America (MOCKFORD 1993). Stenopsocus lachlani Kolbe, 1880 Folicolous species living on conifers. Subdominant species within the whole studied area. Distribution: western Palearctic, missing in the Mediterranean. Elipsocidae Cuneopalpus cyanops (Rostock, 1876) Folicolous species mostly living on coniferous trees, less frequently on deciduous ones. In the PLA Žďárské vrchy only 14 imagoes were caught in the only locality Herálec (HER). In Europe it has only one generation yearly. It was recorded at an altitude of as much as 2,000 m. Distribution: Palearctic. Elipsocus pumilis (Hagen, 1861) Species recorded in 12 localities. Subdominant species within the whole studied area. Distribution: western Palearctic (LIENHARD 1998), probably introduced also into Northern America (MOCKFORD 1993). 215
Hlinsko
HL2
HL1 HLA KAM
Poli�ka
Svratka
dírec nad Doubravou
HER
Krucemburk
KR2
Herálec
STR
KR1
DSK
RBB
MIL
ZKH
CKJ
KJV
Sn�né
CKA
Havlí�kova Borová
JIM
KDL
JIL
Jimramov
STV ZSA
�ár nad Sázavou
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
NMM
Nové M�sto na Morav�
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
49
49
50
50
51
51
52
52
53
53
54
54
55
55
56
56
57
57
58
58
59
59
60
60
61
61
62
62
63
63
64
64
65
65
66
66
67
67
68
68
69
69
70
70
71
71
72
72
73
73
74
74 38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
Fig. 5. Map of researched part of the Protected Landscape Area Žďárské vrchy and particular places of collection (marked by abbreviations)
Reuterella helvimaculata (Enderlein, 1901) Corticolous species found in 5 localities. Imagoes usually live on trunks and branches heavily overgrown with algae and lichens. Subdominant species within the whole studied area. Distribution: western Palearctic. Philotarsidae Philotarsus picicornis (Fabricius, 1793) Corticolous species living on various woody species. It has one or two generations per year in Central Europe. In the PLA Žďárské vrchy caught predominantly on spruce, few individuals also on pine. Eudominant species within the whole studied area. Distribution: Holarctic. 216
Philotarsus parviceps Roesler, 1954 Corticolous species occurring on deciduous and coniferous trees. Dominant species within the whole studied area. In the studied area quite frequently collected together with Philotarsus picicornis. Distribution: Holarctic. HOLUŠA and MÜCKSTEIN (in prep.) reported it as a new species for the Czech Republic. Mesopsocidae Mesopsocus laticeps (Kolbe, 1880) Corticolous species living on deciduous and coniferous trees. It has one generation yearly in Central Europe. In the PLA Žďárské vrchy collected mainly on solitary vegetation (Pinus sylvestris, Alnus glutionosa). Recedent J. FOR. SCI., 49, 2003 (5): 208–219
species within the whole studied area. Distribution: Holarctic. Mesopsocus unipunctatus (Müller, 1764) Corticolous species living on deciduous and coniferous trees. It has one generation per year in Central Europe, overwintering eggs. It was collected in most localities in the PLA Žďárské vrchy, however mainly in monocultures of Picea abies. Dominant species within the whole studied area. Distribution: Holarctic. Psocidae Amphigerontia bifasciata (Latreille, 1799) Corticolous species living on deciduous and coniferous trees. In Central Europe it has one or two generations per year. In the PLA Žďárské vrchy collected only in spruce stands. Subdominant species within the whole studied area. Distribution: Holarctic. Metylophorus nebulosus (Stephens, 1836) Corticolous species living on deciduous and coniferous trees. In Central Europe it has only one generation per year. In the PLA Žďárské vrchy collected mainly in monocultures of Picea abies, also on solitary vegetation (Pinus sylvestris, Alnus glutinosa). Dominant species within the whole studied area. Distribution: Palearctic. Psococerastis gibbosa (Sulzer, 1776) Corticolous species living on deciduous trees, less frequently on coniferous ones. Our largest psocid. It has one generation per year in Central Europe. In the PLA Žďárské vrchy this species was collected sporadically on solitary vegetation, only once it was also found in the undergrowth of fir-beech forest in the locality Ranský Babylon (RBB). Subrecedent species within the whole studied area. Distribution: western Palearctic. Loensia fasciata (Fabricius, 1787) Corticolous species living on deciduous and coniferous tree species that are overgrown with algae and lichens. Imagoes occur already at the beginning of May. The species has one generation yearly, the nymphs of the 3rd instar overwinter. Subrecedent species within the whole studied area. Distribution: western Palearctic. Loensia variegata (Latreille, 1799) Corticolous species living on deciduous and coniferous tree species. It has one generation yearly in Europe. Generally it is rarer than the previous species. Subrecedent species found only in 5 localities within the whole studied area. Distribution: western Palearctic. Trichadenotecnum majus (Kolbe, 1880) Corticolous species living on deciduous and coniferous trees but it always prefers shaded places with higher relative humidity. It has one or two (in climatically favourable years) generations per year. Eudominant species within the whole studied area. Distribution: Holarctic. References
ELLENBERG H. (ed.), 1973. Ökosystemforschung. Berlin – Heidelberg – New York, Springer Verlag: 282. ELLENBERG H., 1978. Vegetation Mitteleuropas mit dem Alpen in ökologischer Sicht. Stuttgart, Eugen Ulmer: 943. GÜNTHER K.K., 1974. Die Tierwelt Deutschlands. 61. Teil. Staubläse, Psocoptera. Jena, VEB Gustav Fischer: 314. HOLUŠA O., 2001. Příspěvek k poznání fauny pisivek (Insecta: Psocoptera) Přírodní rezervace Smrk (Beskydský bioregion, Česká republika). Práce a Stud. Muz. Beskyd, 11: 83–97. HOLUŠA O., MÜCKSTEIN P. (in prep.). Two new species of Psocoptera (Insecta) in the Czech Republic and notes to their ecology. Práce a Stud. Muz. Beskyd. KOLEKTIV, 1994. Lesnický naučný slovník. I. díl A–O. Praha, Ministerstvo zemědělství: 743. KOLEKTIV, 1995. Lesnický naučný slovník. II. díl P–Ž. Praha, Ministerstvo zemědělství: 683. LIENHARD C., 1977. Die Psocopteren des Schweizerischen Nationalparks und seiner Umgebung (Insecta: Psocoptera). Ergebnisse der wissenschaftlichen Untersuchungen im Schweizerischen Nationalpark, Band 14 (Nr. 75): 417–551. LIENHARD C., 1998. Faune de France. Vol. 83. Psocoptéres Euro-méditerranéens. Fédération Française des Sociétes de Sciences naturelles, Paris: 517. MOCKFORD E.L., 1993. Flora & Fauna Handbook No. 10. North American Psocoptera (Insecta). Florida, Gainesville, Sandhill Crane Press: 455. OBR S., 1948. K poznání moravských pisivek. Spisy Přírodov. fakulty MU v Brně, řada M 2, č. 306: 1–107. OBR S., 1949. Pisivky ze Slezských Jeseníků a Králického Sněžníku. Přírodov. Sbor. ostrav. kraje, 10, č. 3: 219–234. OBR S., 1951a. Druhý příspěvek k poznání moravských pisivek. Sbor. Klubu Přírodověd. v Brně, 29: 105–119. OBR S., 1951b. Pisivky Slovenska: I. Vysoké Tatry, II. Okolí Čilistova u Bratislavy. Spisy Přírodov. fakulty MU v Brně, řada M 4, č. 330: 209–229. OBR S., 1952. Pisivky ze Slezských Beskyd. Přírodov. Sbor. ostrav. kraje, 13: 216–231. OBR S., 1958. Třetí příspěvek k poznání moravských pisivek. Sbor. Klubu Přírodověd. v Brně, 30: 53–61. OBR S., 1959. Řád Pisivky – Psocoptera. In: KRATOCHVÍL J. (ed.), Klíč zvířeny ČSR III. Praha, ČSAV: 871. OBR S., 1965. Pisivky Moravských Beskyd. Spisy Přírodov. fakulty UJEP Brno, Serie M 21, L 24, 1965/2 (č. 460): 51–80. PLÍVA K., 1971. Typologický systém ÚHÚL. Brandýs nad Labem, ÚHÚL: 90. PLÍVA K., 1991. Funkčně integrované lesní hospodářství. 1 – Přírodní podmínky v lesním plánování. Brandýs nad Labem, ÚHÚL: 263. PRUNER L., MÍKA P., 1996. Seznam obcí a jejich částí v České republice s čísly mapových polí pro síťové mapování fauny. Klapalekiana, 32 (Suppl.): 1–175. QUITT E., 1975. Klimatické oblasti ČSR 1 : 500 000. Brno, Geografický ústav ČSAV: 1 mapa. Received for publication January 9, 2003 Accepted after corrections February 15, 2003
CULEK M. (ed.), 1996. Biogeografické členění České republiky. Praha, Enigma: 347.
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Složení taxocenóz pisivek (Psocoptera) v závislosti na stupni přirozenosti lesních geobiocenóz v oblasti Žďárských vrchů P. MÜCKSTEIN1, O. HOLUŠA2 Masarykova univerzita, Přírodovědecká fakulta, Brno, Česká republika
1
Mendelova zemědělská a lesnická univerzita, Lesnická a dřevařská fakulta, Brno, Česká republika
2
ABSTRAKT: Na území Chráněné krajinné oblasti Žďárské vrchy byl během let 1999–2000 na vybraných lokalitách studován výskyt pisivek. Celkově bylo uloveno 10 560 imag ve 20 druzích. Materiál byl získáván v různých typech biotopů: přirozené lesní geobiocenózy – porosty buku lesního (Fagus sylvatica) s příměsí jedle bělokoré (Abies alba), javoru klenu (Acer pseudoplatanus) a smrku obecného (Picea abies), přírodě vzdálené geobiocenózy – monokultury smrku obecného (Picea abies), mladé kultury s pasečnou vegetací, rozptýlené soliterní stromy, louky a agrocenózy. Clusterovými analýzami byly vylišeny tři skupiny biotopů: 1. přirozené lesní porosty s dominancí buku lesního, 2. rozptýlená stromová vegetace s keři a jednotlivé soliterní stromy v kulturní krajině, 3. přírodě vzdálené lesní porosty (monokultury smrku obecného). Pro jednotlivé typy biotopů a celkově pro celou zájmovou oblast byla zjištěna dominance jednotlivých druhů. Klíčová slova: pisivky; stupeň přirozenosti lesních geobiocenóz; CHKO Žďárské vrchy; Žďárský bioregion; Česká republika
Na vybraných lokalitách v Chráněné krajinné oblasti Žďárské vrchy byly během vegetační sezony v letech 1999–2000 v pětidenních intervalech sbírány pisivky (Psocoptera). Zkoumaná oblast se nachází v rozpětí nadmořských výšek 583–828 m, tzn. že celá oblast spadá do 5. (jedlo-bukového) vegetačního stupně. Jednotlivé lokality byly voleny tak, aby byly podchyceny taxocenózy pisivek v různých typech biotopů (tab. 1). Celkově byly získáno 10 560 imag ve 20 druzích volně žijících pisivek podřádu Psocomorpha, náležejících do šesti čeledí. Z celkového počtu druhů podřádu Psocomorpha žijících v České republice je to asi 37 %. První imaga byla zjištěna již v první dekádě měsíce května (Loensia fasciata), poslední imaga byla ulovena v polovině listopadu (Caecilius burmeisteri). Maximum početnosti v obou letech bylo zjištěno v měsíci září (obr. 1). Po provedených shlukových analýzách bylo rozděleno 16 studovaných lokalit do tří vzájemně se odlišujících skupin (obr. 2–4): 1. společenstva pisivek žijící v lesních porostech s dominancí buku lesního (Fagus sylvatica); 2. společenstva osídlující skupinky dřevin a keřů a soliterní stromy v otevřené kulturní krajině; 3. společenstva pisivek v monokulturách smrku obecného (Picea abies). V lesních porostech s dominancí buku lesního bylo zjištěno 10 species: eudominantní druhy byly Caecilius flavidus, Philotarsus picicornis, Loensia fasciata, Stenopsocus lachlani, Caecilius burmeisteri; v monokulturách smrku obecného bylo zjištěno 16 druhů: eudominantní – Philotarsus picicornis, Caecilius burmeisteri, Metylophorus nebulosus; v biotopu rozptýlené stromové vegetace a jednotlivých soliterů bylo zjištěno devět druhů: eudominantní – Caecilius flavidus, Loensia fasciata, Mesopsocus unipunctatus (tab. 2). 218
Byly zjištěny dominance druhů pro celkovou zájmovou oblast: eudominantní druhy – Philotarsus picicornis, Caecilius burmeisteri, C. flavidus, dominantní – Loensia fasciata, Mesopsocus unipunctatus, Metylophorus nebulosus, Trichadenotecnum majus, Elipsocus pumilis, Philotarsus parviceps a subdominantní – Caecilius despaxi, Stenopsocus lachlani, Caecilius piceus, Amphigerontia bifasciata, Graphopsocus cruciatus. Hodnoty Shannon-Wienerova indexu diverzity se pohybovaly od 2,13 do 3,48, hodnoty ekvitability se pohybovaly od 0,75 do 0,94. U lokalit se stupněm přirozenosti 2 byly zjištěny hodnoty 2,31–2,62 (ekvitabilita 0,82–0,83), u lokalit se stupněm 4 se hodnoty pohybovaly v rozmezí 2,13–3,25 (ekvitabilita 0,75–0,94) a u lokalit se stupněm 5 se hodnoty pohybovaly v rozmezí 2,24–3,48 (ekvitabilita 0,81–0,93) (tab. 2). Na základě zjištěných výsledků lze konstatovat, že z 54 druhů podřádu Psocomorpha, známých z území ČR, se v CHKO Žďárské vrchy vyskytuje 20 druhů. Tato skutečnost bude zřejmě ovlivněna poměrně vysokou nadmořskou výškou sledovaných lokalit, dosti chladným klimatem a skladbou vegetace (na většině území převládá smrková monokultura). Vyskytují se zde druhy typické jak pro vyšší polohy (Metylophorus nebulosus, Trichadenotecnum majus, Amphigerontia bifasciata), tak druhy žijící spíše v nižších nadmořských výškách (Graphopsocus cruciatus, Caecilius flavidus). Obecně lze říci, že oblast Žďárských vrchů obývají nejběžnější a nejodolnější druhy pisivek s větší ekologickou valencí vzhledem k nadmořské výšce. Z výsledků vyplývá, že taxocenózy pisivek jsou dány kromě klimatických faktorů také porostním typem lesních geobiocenóz. Původní druhové složení se dosti výrazně odlišuje od druhového složení změněných lesních porostů. J. FOR. SCI., 49, 2003 (5): 208–219
Z faunistického a zoogeografického hlediska výzkum přinesl další poznatky o rozšíření druhu Philotarsus parviceps, který je novým druhem pro Českou republiku
(HOLUŠA, MÜCKSTEIN). Tento druh byl ve studované oblasti nalézán poměrně hojně, často společně s druhem Philotarsus picicornis.
Corresponding author: Ing. OTAKAR HOLUŠA, Bruzovská 420, 738 01 Frýdek-Místek, Česká republika tel.: + 420 558 606 690, fax: + 420 558 628 647, e-mail:
[email protected]
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